Specific to papers included in Developmental
Systems Theory (DST) is the belief that the study of development requires a systems-level
model. Such a model would abstract away from the specific biological details of
any particular developmental process in order to isolate the general properties
of developing systems. Contrasting with Developmental
Modularity, DST maintains that identifying the function of individual developmental
modules at the cellular and molecular levels is intractably complicated and is incapable
of representing the structure found at the abstract systems-level, systems
properties are emergent. However, reflecting an internal dispute, the systems
studied are either individual developing organisms (expressing particular
phenotypes) or systems of ecologically-coupled populations of developing organisms
(as they co-evolve with each other).

According to the proponents of Developmental Systems Theory and the Causal Parity Thesis, the privileging of the genome as “first among equals” with respect to the development of phenotypic traits is more a reflection of our own heuristic prejudice than of ontology - the underlying causal structures responsible for that specified development no more single out the genome as primary than they do other broadly “environmental” factors. Parting with the methodology of the popular responses to the Thesis, this paper offers (...) a novel criterion for ‘causal primacy’, one that is grounded in the ontology of the unique causal role of dispositional properties. This paper argues that, if the genome is conceptualised as realising dispositional properties that are “directed toward” phenotypic traits, the parity of ‘causal roles’ between genetic and extra-genetic factors is no longer apparent, and further, that the causal primacy of the genome is both plausible and defensible. (shrink)

We show how a simple nonlinear dynamical system (the discrete quadratic iteration on the unit segment) can be the basis for modelling the embryogenesis process. Such an approach, even though being crude, can nevertheless prove to be useful when looking with the two main involved processes:i) on one hand the cell proliferation under successive divisions ii) on the other hand, the differentiation between cell lineages. We illustrate this new approach in the case of Caenorhabditis elegans by looking at the early (...) stages of embryogenesis, up to several hundreds of cells (lima bean larval stage). We show how the many results that have been obtained by several groups can be interpreted in terms of values for the parameters controlling the dynamical system. Furthermore, we can extend the model to the cases of genetic mutations. More precisely the teratogenetic and lethal effects are associated with abnormal variation of the control parameters with time. (shrink)

Possible effects of interaction (cross-talk) between signaling pathways is studied in a system of Reaction–Diffusion (RD) equations. Furthermore, the relevance of spontaneous neurite symmetry breaking and Turing instability has been examined through numerical simulations. The interaction between Retinoic Acid (RA) and Notch signaling pathways is considered as a perturbation to RD system of axon-forming potential for N2a neuroblastoma cells. The present work suggests that large increases to the level of RA–Notch interaction can possibly have substantial impacts on neurite outgrowth and (...) on the process of axon formation. This can be observed by the numerical study of the homogeneous system showing that in the absence of RA–Notch interaction the unperturbed homogeneous system may exhibit different saddle-node bifurcations that are robust under small perturbations by low levels of RA–Notch interactions, while large increases in the level of RA–Notch interaction result in a number of transitions of saddle-node bifurcations into Hopf bifurcations. It is speculated that near a Hopf bifurcation, the regulations between the positive and negative feedbacks change in such a way that spontaneous symmetry breaking takes place only when transport of activated Notch protein takes place at a faster rate. (shrink)

The meaning of the concept of fusion is discussed in relation with the works of Payer and those of Van Tieghem. It is pointed out that there is a difference, at the theoretical level, between the concept of fusion congénitale as defined by Payer and the concept of concrescence congénitale formulated by Van Tieghem. The former is inobservable by definition, while the latter deals with intercalary growth. For Van Tieghem, anatomy can prove the existence of fusion, even if we do (...) not see it during ontogenesis.We distinguish three complementary methods for explaining the unions of organs: ontogenetic, typological and phylogenetic. We have attempted, not so much to defend one or other of these methods, as to show that they often invoke very different interpretations for the same morphological phenomena. It is probable that only an analysis of the writings of 19th century botanists will clarify the concept of fusion and more generally the epistemology of plant morphology. (shrink)

The Developmental Systems Theory (DST) presented by its proponents as a challenging approach in biology is aimed at transforming the workings of the life sciences from both a theoretical and experimental point of view (see, in particular, Oyama [1985] 2000; Oyama et al. 2001). Even though some may have the impression that the enthusiasm surrounding DST has faded in very recent years, some of the key concepts, ideas, and visions of DST have in fact pervaded biology and philosophy of biology. (...) It seems crucial to us both to establish which of these ideas are truly specific to DST, and to sift through these ideas in order to determine the criticisms they have drawn, or may draw (e.g., Sterelny et al. 1996; Griesemer 2000; Sterelny 2000; Kitcher 2001; Keller 2005; Waters 2007). (shrink)

This paper juxtaposes Deleuze's notion of the virtual alongside Oyama's notion of a developmental system in order to explore the promises and perils of thinking bodily identity as indeterminate at a time when new technologies render bodily ambiguity increasingly productive of both economic profit and power relations.

Comparing engineering to evolution typically involves adaptationist thinking, where well-designed artifacts are likened to well-adapted organisms, and the process of evolution is likened to the process of design. A quite different comparison is made when biologists focus on evolvability instead of adaptationism. Here, the idea is that complex integrated systems, whether evolved or engineered, share universal principles that affect the way they change over time. This shift from adaptationism to evolvability is a significant move for, as I argue, we can (...) make sense of these universal principles without making any adaptationism claims. Furthermore, evolvability highlights important aspects of engineering that are ignored in the adaptationist debates. I introduce some novel engineering examples that incorporate these key neglected aspects, and use these examples to challenge some commonly cited contrasts between engineering and evolution, and to highlight some novel resemblances that have gone unnoticed. (shrink)

Like Laland et al., I think Mayr’s distinction is problematic, but I identify a further problem with it. I argue that Mayr’s distinction is a false dichotomy, and obscures an important question about evolutionary change. I show how this question, once revealed, sheds light on some debates in evo-devo that Laland et al.’s analysis cannot, and suggest that it provides a different view about how future integration between biological disciplines might proceed.

John Dupr explores recent revolutionary developments in biology and considers their relevance for our understanding of human nature and society. He reveals how the advance of genetic science is changing our view of the constituents of life, and shows how an understanding of microbiology will overturn standard assumptions about the living world.

Among the factors necessary for the occurrence of some event, which of these are selectively highlighted in its explanation and labeled as causes — and which are explanatorily omitted, or relegated to the status of background conditions? Following J. S. Mill, most have thought that only a pragmatic answer to this question was possible. In this paper I suggest we understand this ‘causal selection problem’ in causal-explanatory terms, and propose that explanatory trade-offs between abstraction and stability can provide a principled (...) solution to it. After sketching that solution, it is applied to a few biological examples, including to a debate concerning the ‘causal democracy’ of organismal development, with an anti-democratic (though not a gene-centric) moral. (shrink)

Freedom is first apprehended as the pursuit of an activity which implies the choice to defend a thesis among other possible ones. This translation of the problem of freedom in an articulate language presupposes a complex nervous system and sensory apparatuses which we take for granted. In this study, I try to explore the undergrounds of the problem of freedom along with the suggestion that the notion of coding could enable one to bridge nature and the mind. When organisms invent, (...) are they doing it in a spontaneous manner, inscribing in their hereditary and mnemonic instructions a stochastic contrivance of random accidents, or are they attempting to select among a limited number of schemes endowed with some optimality of functioning? If we consider them as submitted to physical forces, it is to the extent that we make them part of a strategy to extend the "laws" of a nature understood to respond passively. I suggest in this study that the epistemological understanding must regionalize itself and admit a hierarchy of dispositions in relation to the phenomenon of selection. I end by suggesting the pursuit of affirmation instead of negation, as this alone contains the requirement of integration to the knowing subject as well as the form in its act of understanding, without giving it a spontaneist position. (shrink)

Some central ideas associated with developmental systems theory (DST) are outlined for non-specialists. These ideas concern the nature of biological development, the alleged distinction between "genetic" and "environmental" traits, the relations between organism and environment, and evolutionary processes. I also discuss some criticisms of the DST approach.

Developmental systems theory (DST) is a general theoretical perspective on development, heredity and evolution. It is intended to facilitate the study of interactions between the many factors that influence development without reviving `dichotomous' debates over nature or nurture, gene or environment, biology or culture. Several recent papers have addressed the relationship between DST and the thriving new discipline of evolutionary developmental biology (EDB). The contributions to this literature by evolutionary developmental biologists contain three important misunderstandings of DST.

The Developmental Systems approach to evolution is defended against the alternative extended replicator approach of Sterelny, Smith and Dickison (1996). A precise definition is provided of the spatial and temporal boundaries of the life-cycle that DST claims is the unit of evolution. Pacé Sterelny et al., the extended replicator theory is not a bulwark against excessive holism. Everything which DST claims is replicated in evolution can be shown to be an extended replicator on Sterelny et al.s definition. Reasons are given (...) for scepticism about the heuristic value claimed for the extended replicator concept. For every competitive, individualistic insight the replicator theorist has a cooperative, systematic blindspot. (shrink)

In 1990 Robert Lickliter and Thomas Berry identified the phylogeny fallacy, an empirically untenable dichotomy between proximate and evolutionary causation, which locates proximate causes in the decoding of ‘genetic programs’, and evolutionary causes in the historical events that shaped these programs. More recently, Lickliter and Hunter Honeycutt (Psychol Bull 129:819–835, 2003a) argued that Evolutionary Psychologists commit this fallacy, and they proposed an alternative research program for evolutionary psychology. For these authors the phylogeny fallacy is the proximate/evolutionary distinction itself, which they (...) argue constitutes a misunderstanding of development, and its role in the evolutionary process. In this article I argue that the phylogeny fallacy should be relocated to an error of reasoning that this causal framework sustains: the conflation of proximate and evolutionary explanation. Having identified this empirically neutral form of the phylogeny fallacy, I identify its mirror image, the ontogeny fallacy. Through the lens of these fallacies I attempt to solve several outstanding problems in the debate that ensued from Lickliter and Honeycutt’s provocative article. (shrink)

Will a synthesis of developmental and evolutionary biology require a focus on the role of nongenetic resources in evolution? Nongenetic variation may exist but be hidden because the phenotypes are stable (developmentally canalized) under certain background conditions. In this case, those differences may come to play important roles in evolution when background conditions change. If this is so, then a focus on the way that developmental resources are made reliable, and the ways in which reliability fails, may prove to be (...) of crucial importance to linking developmental and evolutionary biology. †To contact the author, please write to: 208 Hovland Hall, Philosophy Department, Oregon State University, Corvallis, OR 97331‐3902; e‐mail: jonathan.kaplan@oregonstate.edu. (shrink)

In 1961, Ernst Mayr published a highly influential article on the nature of causation in biology, in which he distinguished between proximate and ultimate causes. Mayr argued that proximate causes (e.g. physiological factors) and ultimate causes (e.g. natural selection) addressed distinct ‘how’ and ‘why’ questions and were not competing alternatives. That distinction retains explanatory value today. However, the adoption of Mayr’s heuristic led to the widespread belief that ontogenetic processes are irrelevant to evolutionary questions, a belief that has (1) hindered (...) progress within evolutionary biology, (2) forged divisions between evolutionary biology and adjacent disciplines and (3) obstructed several contemporary debates in biology. Here we expand on our earlier (Laland et al. in Science 334:1512–1516, 2011) argument that Mayr’s dichotomous formulation has now run its useful course, and that evolutionary biology would be better served by a concept of reciprocal causation, in which causation is perceived to cycle through biological systems recursively. We further suggest that a newer evolutionary synthesis is unlikely to emerge without this change in thinking about causation. (shrink)

Recent and not so recent advances in our molecular understanding of the genome make the once prevalent view of the genome as a passive container of genetic information (i.e., genes) untenable, and emphasize the importance of the internal organization and re-organization dynamics of the genome for both development and evolution. While this conclusion is by now well accepted, the construction of a comprehensive conceptual framework for studying the genome as a dynamic system, capable of self-organization and adaptive behavior is still (...) underway. This work deals with the effect of such a conceptual shift on evolutionary thought. Specifically, I try to articulate the conceptual commitments and obligations of views that privilege explanatorily or causally the genome, its dynamics and mechanisms, over genes. I refer to this class of views as belonging to ‘the genome perspective’. (shrink)

In reworking a variety of biological concepts, Developmental Systems Theory (DST) has made frequent use of parity of reasoning. We have done this to show, for instance, that factors that have similar sorts of impact on a developing organism tend nevertheless to be invested with quite different causal importance. We have made similar arguments about evolutionary processes. Together, these analyses have allowed DST not only to cut through some age-old muddles about the nature of development, but also to effect a (...) long-delayed reintegration of development into evolutionary theory. Our penchant for causal symmetry, however (or 'causal democracy', as it has recently been termed), has sometimes been misunderstood. This paper shows that causal symmetry is neither a platitude about multiple influences nor a denial of useful distinctions, but a powerful way of exposing hidden assumptions and opening up traditional formulations to fruitful change. (shrink)

The Modern Synthesis of Darwinism and genetics regards non-genetic factors as merely constraints on the genetic variations that result in the characteristics of organisms. Even though the environment (including social interactions and culture) is as necessary as genes in terms of selection and inheritance, it does not contain the information that controls the development of the traits. S. Oyama’s account of the Parity Thesis, however, states that one cannot conceivably distinguish in a meaningful way between nature-based (i.e., gene-based) and nurture-based (...) (i.e., environment-based) characteristics in development because the information necessary for the resulting characteristics is contained at both levels. Oyama and others argue that the Parity Thesis has far-reaching implications for developmental psychology, in that both nativist and interactionist developmental accounts of motor, cognitive, affective, social, and linguistic capacities that presuppose a substantial nature/nurture dichotomy are inadequate. After considering these arguments, we conclude that either Oyama’s version of the Parity Thesis does not differ from the version advocated by liberal interactionists, or it renders precarious any analysis involving abilities present at birth (despite her claim to the contrary). More importantly, developmental psychologists need not discard the distinction between innate characteristics present at birth and those acquired by learning, even if they abandon genocentrism. Furthermore, we suggest a way nativists can disentangle the concept of maturation from a genocentric view of biological nature. More specifically, we suggest they can invoke the maturational segment of the developmental process (which involves genetic, epigenetic and environmental causes) that results in the biological “machinery” (e.g. language acquisition device) which is necessary for learning as a subsequent segment of the developmental process. (shrink)

In a now classic paper published in 1991, Alberch introduced the concept of genotype–phenotype (G!P) mapping to provide a framework for a more sophisticated discussion of the integration between genetics and developmental biology that was then available. The advent of evo-devo first and of the genomic era later would seem to have superseded talk of transitions in phenotypic space and the like, central to Alberch’s approach. On the contrary, this paper shows that recent empirical and theoretical advances have only sharpened (...) the need for a different conceptual treat- ment of how phenotypes are produced. Old-fashioned metaphors like genetic blueprint and genetic programme are not only woefully inadequate but positively misleading about the nature of G!P, and are being replaced by an algorithmic approach emerging from the study of a variety of actual G!P maps. These include RNA folding, protein function and the study of evolvable soft- ware. Some generalities are emerging from these disparate fields of analysis, and I suggest that the concept of ‘developmental encoding’ (as opposed to the classical one of genetic encoding) provides a promising computational–theoretical underpinning to coherently integrate ideas on evolvability, modularity and robustness and foster a fruitful framing of the G!P mapping problem. (shrink)

In the fall of 1990 I had just began my doc- toral studies at the University of Connecticut. Freshly arrived from Italy, I came to the United States to work with Carl Schlichting on something to do with phenotypic plastic- ity. I spent most of that semester discussing with other graduate students what I thought was a momentous paper by Mary Jane West- Eberhard (1989) in the Annual Review of Ecol- ogy and Systematics. That paper, entitled Phe- notypic Plasticity and (...) the Origins of Diversity, was a (quite lengthy) forerunner of the (also quite bulky) book I am reviewing now. Like the paper, this volume has the potential to be momentous in the development of our ideas on phenotypic evolution. (shrink)

In this paper, I address the question of what the Developmental Systems Theory (DST) aims at explaining. I distinguish two lines of thought in DST, one which deals specifically with development, and tries to explain the development of the individual organism, and the other which presents itself as a reconceptualization of evolution, and tries to explain the evolution of populations of developmental systems (organism-environment units). I emphasize that, despite the claiming of the contrary by DST proponents, there are two very (...) different definitions of the ‘developmental system’, and therefore DST is not a unified theory of evolution and development. I show that the DST loses the most interesting aspects of its reconceptualization of development when it tries to reconceptualize evolutionary theory. I suggest that DST is about development per se, and that it fails at offering a new view on evolution. (shrink)

How does a complex organism develop from a relatively simple, homogeneous mass? The usual answer is: through the execution of species-specific genetic instructions specifying the development of that organism. Commentators are sometimes sceptical of this usual answer, but of course not all commentators. Some biologists refer to master control genes responsible for the activation of all the genes responsible for every aspect of organismal development; and some philosophers, most notoriously Rosenberg, buy this claim hook, line, and sinker. Here I explore (...) both the seeming plausibility of the usual position, and also its ultimate inadequacy. (shrink)

The question which is never entirely resolved is: what is life? Biology, claims to stand for the study of life and living things, yet we would say that it cannot make a thoroughly clear distinction between living and non living, except in some very obvious cases. There are textbook definitions, of course, based on certain notable properties such as the ability to metabolize or reproduce, but these are arbitrary. If we are familiar with the characteristics of a particular animal or (...) plant, we know enough to be able to pronounce that it is dead when certain internal and external behaviours are no longer evident. Even this has difficulties - such as revealed in the arguments about whether to switch off a human life support system or not. When you find a squishy object on the seashore, can you be sure if it is alive or dead - or never living? The same dilemma confronts medical scientists and microbiologists trying to decide, for example, whether viruses are living, or quasi living, or intermittently living, or what. (shrink)

I critically assess two widely cited evolutionary biological arguments for two versions of the ‘Extended Mind Thesis’ (EMT): namely, an argument appealing to Dawkins’s ‘Extended Phenotype Thesis’ (EPT) and an argument appealing to ‘Developmental Systems Theory’ (DST). Specifically, I argue that, firstly, appealing to the EPT is not useful for supporting the EMT (in either version), as it is structured and motivated too differently from the latter to be able to corroborate or elucidate it. Secondly, I extend and defend Rupert’s (...) argument that DST also fails to support or elucidate the EMT (in either version) by showing that the considerations in favour of the former theory have no bearing on the truth of the latter. I conclude by noting that the relevance of this discussion goes beyond the debate surrounding the EMT, as it brings out some of the difficulties of introducing evolutionary biological considerations into debates in psychology and philosophy more generally. (shrink)

Although the construction of neo-Darwinism grew out of Thomas Hunt Morgan's melding of Darwinism and Mendelism, his evidence did not soley support a model of gradual change. To the contrary, he was confronted with observations that could have led him to a more "evo-devo" understanding of the emergence of novel features. Indeed, since Morgan was an embryologist before he became a fruit-fly geneticist, one would have predicted that the combination of these two lines of research would have resulted in early (...) formulations of concepts relevant to evolutionary developmental biology. It is thus of interest to review Morgan's thought processes and arguments for at first rejecting both Darwinism and Mendelism, and then for later dismissing data that would have yielded a model of rapid morphological change in favor of a model of change based on the accumulation of minor mutations and their morphological consequences. (shrink)

Evolution proceeds in phases, alternatingly convergent and divergent. During the divergent phases, many variants of an evolutionary system arise, and in the convergent phases, these are brought together in a new, higher unity, which in turn varies, and so on. Thus the mechanism of evolution is trialistic, proceeding according to the Hegelian principle (in the widest sense) of thesis, antithesis and synthesis. This mechanism is at the same time mirrored in the structure of the evolving systems, being most clearly expressed (...) in the derivation of periodic systems of the individual levels of evolution. These relationships will be discussed using examples from symbiosis research, population dynamics and biogenesis. (shrink)

The concept of innateness is used to make inferences between various better-understood properties, like developmental canalization, evolutionary adaptation, heritability, species-typicality, and so on (‘innateness-related properties’). This article uses a recently-developed account of the representational content carried by inheritance systems like the genome to explain why innateness-related properties cluster together, especially in non-human organisms. Although inferences between innateness-related properties are deductively invalid, and lead to false conclusions in many actual cases, where some aspect of a phenotypic trait develops in reliance on (...) a genetic representation it will tend, better than chance, to have many of the innateness-related properties. The account also shows why inferences between innateness-related properties sometimes fail and argues that such inferences are especially misleading when applied to human psychology and behaviour because human psychological development is especially reliant on non-genetic inherited representations. (shrink)

Developmental Systems Theory (DST) emphasises the importance of non-genetic factors in development and their relevance to evolution. A common, deflationary reaction is that it has long been appreciated that non-genetic factors are causally indispensable. This paper argues that DST can be reformulated to make a more substantive claim: that the special role played by genes is also played by some (but not all) non-genetic resources. That special role is to transmit inherited representations, in the sense of Shea (2007: Biology and (...) Philosophy, 22, 313-331). Formulating DST as the claim that there are non-genetic inherited representations turns it into a striking, empirically-testable hypothesis, driving the sort of investigations that are only now beginning to appear in the scientific literature. DST’s characteristic rejection of a gene vs. environment dichotomy is preserved, but without dissolving all potentially explanatory distinctions into an interactionist causal soup, as some have alleged. (shrink)

A central idea of developmental systems theory is ‘parity’ or ‘symmetry’ between genes and non-genetic factors of development. The precise content of this idea remains controversial, with different authors stressing different aspects and little explicit comparisons among the various interpretations. Here I characterise and assess several influential versions of parity.

In this paper I develop three conceptions of the relationship between evolutionary and developmental biology. I further argue that: (a) the choice between them largely turns on as yet unresolved empirical considerations; (b) none of these conceptions demand a fundamental conceptual reevaluation of evolutionary biology; and (c) while developmental systems theorists have constructed an important and innovative alternative to the standard view of the genotype/phenotype relations, in considering the general issue of the relationship between evolutionary and developmental biology, we can (...) remain neutral on this debate. (shrink)